CATRINA (2008), 3 (1): 1 10 PROCEEDINGS OF THE SECOND INTERNATIONAL CONFERENCE ON THE ROLE OF GENETICS AND © 2008 BY THE EGYPTIAN SOCIETY FOR ENVIRONMENTAL SCIENCES BIOTECHNOLOGY IN CONSERVATION OF NATURAL RESOURCES , ISMAILIA , EGYPT , JULY 9 10, 2007 Systematic Review of the Papaveracea Adans. and Status of Some Genera Wafaa K. Taia Alexandria University, Faculty of Science, Botany Department, Alexandria Egypt ABSTRACT This work deals with the systematic relations of family Papaveraceae according to the new data derived from molecular analysis. An overall view has been given to elucidate the taxonomic position of the family within the basal eudicots. Characteristic features of all the group has been given with clear view of the general features of the family and its taxonomic divisions. Complete list of the genera, as listed by Royal Botanical Gardens in Kew has been added with summary of the recent phylogenetic relationships of the family and its genera. Leaf and fruit morphological variations of 14 species belonging to Papaveraceae s.s., 7 species belonging to Fumariaceae s.s. and two species from Hypecoaceae s.s. have been studied to assess the new classification of the Papaveraceae s.l. with complete description of their morphological variations and their position according to the new classification of the family. Keywords: Eudicots, Magnoliopsida, Papaveraceae, Phylogeny, Ranunculales, Taxonomy. Taxonomic Status of the Papaveraceae s.l. Ranunculaceae , Papaveraceae, Fumariaceae, The status of family Papaveraceae s.l. has been faced Berberidaceae, Menispermaceae , and Aristolochiaceae . with numerous opinions. Before the extensive ribosomal The family is closely related to the Fumariaceae which and DNA sequences researches concerning the often included within it (Norris, 1941, Fohne, 1962, phylogeny of the taxonomic taxa, the family has been Kolbe, 1978, Behnke and Barthlott, 1983 Chase et al., considered by Wettstein (1935) as one of the 1993; Taia and Sheha, 2003). Meanwhile, the family Rhoeadales families. Later, Melchior (1964), Tamura has close affinity to members of Ranunculales with the (1974), Benson (1979) and Tutin et al. (1993) renamed only differences that gynoecium is paracarpous and the the order Rhoeadales by Papaverales s.l. with four presence of secretory idioblasts or laticifers in the suborders; Papaverinae, Capparinae, Tovarianae and Papaveraceae s.l., and both families are considered as Moringanae . The family was found to be closely related primitive families. Parker (1982) separated the to the Brassicaceae , Capparidaceae , Resedaceae , Papaveraceae as an order Papaverales while the Tovariaceae and Moringaceae (Blagowestschenski, Ranunculaceae as order Ranunculales and both orders 1955, Hegnauer, 1961 and Gershenzon and Mabry, under subclass Magnoliidae , class Magnoliopsida . This 1983). The position of these families in the same order class refers to a small group containing Papaveraceae, is due to flower characters, such as regular and Ranunculaceae and Berberidaceae as sister group to hypogynous flowers, anthers arranged in several whorls Monocots. Based on the recent work on molecular and the carpels are syncarpous and possess two to many analysis, done by Hoot et al. (1999), the Papaveraceae parietal ovules. Norris (1941) considered the s.l. is considered as one of the families belonging to Papaveraceae s.l. closely related to both the order Ranunculales and they up graded the order Fumariaceae and Brassicaceae and they all evolved Ranunculales from Magnoliidae to the base of the from the same origin according to torus anatomy and Eudicots, which is in close association with the nectary characteristics. Monocots. Family Papaveraceae Adans. nom.cons. s.l . which is a north temperate, mostly herbaceous family consisting Phylogenetic Opinions within the Family of 23 genera and about 250 species (Heywood, 1993), Cronquist (1981) considered the Ranunculales as one Newly recorded species raises the number of genera to of the woody magnoliids because their flowers have free 35 as compared with the list given by Royal Botanical parts that are sometimes spirally arranged. Cronquist Gardens, in Kew, 2006. The family has three synonyms; (1981) suggested that the connection to the woody Chelidoniaceae Martinov., Echscholziaceae Ser. and magnoliids was via Illiciaceae (ANITA grade; Qui et Platystemonaceae (Rchb.ex Spatch) Lilja. According to al. , 1999) due to the presence of triaperturate pollen Cronquist System (1981), family Papaveraceae s.l. is grains in both groups. Spichiger and Savolainen (1997) classified under subclass Magnoliidae together with the pointed to the similarities between the Ranunculales and Magnoliaceae , Nymphaeaceae and Ranunculaceae . Papaverales with the monocots as they share many This subclass is characterized by well developed features such as imperfect vessels; inaperturate or flowers with separated perianth or calyx and corolla, the uniaperturate pollen grains (or derived types). Recent stamens are numerous and gynoecium is apocarpous. In works based on molecular analysis considered the Papaveraceae gynoecium is paracarpous. According to Ranunculales as a monophyletic group (Fig. 1) (Chase South West Virginia flora, the Magnoliideae comprises et al. , 1993, Drinnan et al., 1994, Hoot and Crane 1995, nine families; Magnoliaceae , Annonaceae, Lauraceae, Soltis et al. , 1997, Kallersjo et al. , 1998, ______ * Corresponding author: [email protected] Systematic review of the Papaveracea and status of some genera
General Features of the Family The family has plants in different varieties of life forms and morphological characters. They are herbs or sub shrubs, shrubs, or even small trees ( Dendromegon rigida) , annuals, biennials, or perennials. Plants have tap roots or rhizomes with leafy or naked erect or spreading stems. Leaves are basal and/or cauline, alternate to opposite or whorled, simple ex stipulate petiolate or sessile with entire blade or lobed in pinnate, subpalmate, or palmate orders of lobes. In Argemone the leaves are dissected with spiny margins. The leaves are mostly glabrous, except few species covered with ramified or glandular hairs with smooth walls. Stomata are either diacytic or anomocytic with isodiametric or elongated epidermal cells (Heywood, 1993). The family has variety of flower arrangements, forms and colors, but all are bracteate, radially symmetric, pedicellate or sessile; receptacle sometimes expanded and forming cup or ring beneath calyx (in Eschscholzia, Figure (1) : Phylogeny of the Ranumculales , showing relative Meconella and Platystemon) . The flowers are either positions of the major families within this order (Tree solitary or arranged in groups which are either terminal adapted from Hoot et al., 1999). Numbers above lines or axillary. The inflorescences are either cymose or indicate the number of nucleotide changes supporting each racemose, umbelliform or corymbiform. Flowers are branch. Number below the branches is the percentage of sometimes perigynous; sepals two or three, ob ovate, times that the branch was recovered in 1000 bootstrap distinct or connate always caducous. Petals are distinct, replications. colored and ob ovate, usually four or more, sometimes absent. Stamens are numerous in many whorls, Hoot et al. , 1999). Hilu et al. (2003), Kim et al. (2004) sometimes 4 15 in Meconella and Canbya , with bi and Worberg et al. (2006) supported the close locular anthers (Kiger, 1996). Pollen grains appear association of the Papaveraceae s.l. with the spheroidal or sub prolate, medium sized, usually with Fumariaceae then with the Eupteleaceae and the three tricolpate apertures. In some species such as Papaver families are in close relation with the rest of the argemone, Argemone mexicana, and Roemeria hybrida , Ranunculales families. pollen grains are polyporate (Taia and Sheha, 2003). Despite the relations between the Papaveraceae s.l. Pistil is 1, 2 to many (22) united carpels with one or genera are still obscure, due to the lack of fossils two locules, sometimes multilocular by placental records for that group and their geographical intrusion, placenta two or more in parietal position. distribution. Kadereit et al. (1994) found that the genera Style is usually one or absent with sessile stigma. Papaver L., Meconopsis Vig., Stylomecon G.Taylor and Stigma lobes are 2 to many in circular disc or radiating Roemeria Medik. Belonging to Papaveraceae s.str. ones. Fruits are capsules dehiscent by pores, valves or subfamily Papaveroideae are morphologically similar. dissociating and breaking transversely into one seeded In the mean time Schwarzbach & Kadereit (1995) segments (only in Platystemon ). Seeds are always many reported that these genera form a monophyletic clade on small, sometimes arillate or carunculate with different the basis of molecular analysis. Kadereit et al. (1997), colors varies from white to black or different shades of on the basis of molecular data concluded that (1) Asian brown. Meconopsis is not a monophyletic group, but paraphyletic in relation to Papaver s.l. including Classification of the Family M.cambrica , Stylomecon and Roemeria ; (2) both The Papaveraceae s.l. is subdivided into three Roemeria and Stylomecon are nested with Papaver s.l., subfamilies by Hoot et al., 1997; Pteridophylloideae, Roemeria is sister group to Papaver sect. Argemonidium Papaveroideae , and Fumarioideae. Ernst (1962), Layka and Stylomecon to papaver californicum ; and (3) the (1976), Heslop Harison and Shivana (1977), Mabry position of M.cambrica within papaver s.l. (incl. (1973), Kaderreit (1993), Kaderreit et al. (1994) and Stylomeco n and Roemeria ) allows the arise of it from Bruckner (2000) divided the family into four within Papaver s.l. in parallel to Asian Meconopsis and subfamilies; Chelidonoideae , Eschscholzioideae , in this case the genus Papave r will be monophyletic – or Papaveroideae and Platystemonoideae . Fumaroideae M. cambrica can be regarded as an original although and Pteridophylloideae were separated in new families, disjunct Meconopsis , in this case the entire genus Fumariaceae , and Pteridophyllaceae , and the four tribes Meconopsis would be paraphyletic in relation to under subfamily Papaveroideae have been graded up to Papaver s.l. – or both Meconopsis and Papaver s.l. are the rank of subfamilies. This division is based mainly on polyphyletic. gynoecium morphology and indumentum's characters
2 Kamal W.T.
but the evolutionary relationships within the subfamilies St., P. hybridum L., P. mexicana L., P. polytrichum remain ambiguous as well as the relations between the Boiss , P. somniferum L. and P. rhoeas L.. While genus genera. Hoot et al. (1999) on the basis of molecular data Glaucium has four species; G. arabicum Fres, G. found that the genera Corydalis and Hypecoum corniculatum L., G. Flavum Crantz, and G. belonging to the Fumarioidae are closely related and grandiflorum Boiss & Huet. The other two genera are they supported their separation in the new family represented only by one species each, Argemone Fumariaceae . The same to the genus Pteridophyllum , is mexicana L. and Roemeria hybridica Medic. while better to be treated as a separate family family Fumariaceae has one genus; Fumaria with eight Pteridophyllaceae , while the rest of the genera are species in Egypt. These are F. bracteosa Pomel, F. better to be subdivided into three subfamilies (Fig. 2). capreolata L., F. densiflora DC., F. gaillardotii Boiss., They are Adlumia Raf ex DC., Arctomecon Torr. & F.judaica Boiss. F. microstachys Kral., F. officinalis L., Frem. Argemone L., Bocconia L., Canbya Parry ex A. and F. parviflora Lam. Moreover genus Hypecoum L. is Gray, Capnoides Mill., Ceratocapnos Durieu, separated in a distinct monogeneric family Hypecoaceae Chelidonium L., Cryptocapno s Rech.F., Cysticapnos with eight species which show relation to the Mill., Dactylicapnos Wall., Dendromecon Benth., Fumariaceae (Täckholm, 1974). Dicentra Borkh. Ex Bemh., Dicranostigma Hook.f. & Thomson, Discocapnos Cham. & Schltdl., Eomecon MATERIALS AND METHODS Hance, Eschscholzia Cham., Glaucium Mill., Herbarium specimens allocated at Alexandria Hesperomecon Greene, Hunnemannia Sweet, University Herbarium (AUH) have been examined Hylomecon Maxim., Hypecoum L., Macleaya R.Br., carefully to clarify leaf and fruit morphological Meconella Nutt., Meconopsis R. Vig., Papave r L., variations within the genera. List of the examined Platycapnos (DC.) Bernh., Platystemon Benth., specimens, their collectors and localities are given in Pteridophyllum Siebold & Zucc., Roemeria Medik., table 1. The studied taxa include some species from Romneya Harv., Rupicapnos Pomel, Sanguinaria L., USA and France and represent seven genera and 23 Stylomecon G.Taylor, Stylophorum Nutt. And species . Leaf and fruit morphological measurements (in Trigonocapnos Schltr. Some of these genera contain one cm.) and descriptions are summarized in table (2). species only which is endemic to certain localities, Photographs of selected species to clarify the while others like Papaver, Argemone, Glaucium, differences in leaf and fruit characters have been also Meconella, Meconopsis, Eschscholzia, Dendromegon, given (plate 1). Chelidonium and Roemeria are more abundant and well known. RESULTS AND DISCUSSION The careful examinations of the herbarium sheets Family Papaveraceae showed that the three families; Papaveraceae, Fumariaceae and Hypecoaceae , are closely related. In Subfamily Fumarioideae all the studied taxa, the leaves are simple, radical or cauline, petiolated with long petioles except in few Tribe Hypecoeae species; G. arabicum, R. hybrida, F. microstachys, and Tribe Fumarieae H. pendulum ; the leaves are sessile. The genera Fumaria, Dicentra and Hypecoum have glabrous plants, Subfamily Papaveroideae while the genera Roemeria, Papaver, Glaucium and Eschscholzia plants are covered by long multicellular, Tribe Papavereae uniserriate hairs and the fruits are supported, as well, by Tribe Chelidoneae long hairs. The leaves are crowded at the base of the plants even in plants with cauline leaves. In Eshscholzia Tribe Eschscholtzieae species, the leaf blades are either very finely dissected or finely dissected and appears as being in tufts. The Tribe Platystemoneae genera Glaucium, Papaver and Dicentra have pinnatifid Subfamily Pteridophylloideae or slightly lacerus leaf blades. Species of Fumaria have alternate or laxus; soft and pendulous without fixed Figure (2) : Family Papaveraceae Classification according to arrangement; leaves and very finely dissected leaf Hoot et al. (1997). blades. Stigma and fruit shapes are very important in the According to Täckholm (1974) , the Egyptian differentiation of the species, as all the Eschscholzia and Papavercaceae s.s. comprises four native genera Roemeria species have pointed stigmas, with very short Papaver, Argemone, Roemeria and Glaucium and one style and linear fruits. Glaucium species have capitate, introduced genus; Eschscholzia . These genera are rounded stigmas with very short style and linear fruits. narrowly distributed, in Egypt they are mainly In the three genera the fruits are siliqua, but Mediterranean annual species. Genus Papaver has Eschscholzia has glabrous ones while the latter two seven species: P. argemone L., P. decaisnei Hochst & have hairy fruits. Papaver species have capitate, sessile
3 Systematic review of the Papaveracea and status of some genera
Table (1): Studied taxa and their collectors and localities.
TAXA COLLECTOR LOCALITY Papaveraceae s.s. 1-Eschscholzia E. caespitosa Benth. Peta C. Lewis 4 ( 16/4/1967) Napa County, California E. californica Cham Linda Miller 11 (16/4/1967) Solano County, California Loise Krozek 12 (22/4/1965) Yolo County, California Jean Addicott 16 Marin County, California E. glyptosperma Greene June McCaskill 280 (30/4/1954) Inyo County, California E. lemmonii Greene J.M. Tucker 2351 (19/4/1952) San Luis Obispo County, California E. lobii Greene Anne Harrison 15 (2/4/1962) Solano County, California E. minutifolia S. Wats. E.M. Gifford (27/4/1950) Inyo County, California J. McCaskill 241 (29/4/1954) Inyo County, California 2-Glaucium G. arabicum Fresen L. Boulos (24/6/2005) Southern Sinai, St.Katherine, Egypt G. corniculatum (L.)J.H. S. Kamal (4/5/2003) Khorshid, Egypt S. Rashad (19/3/1948) King Mariut, Egypt S. Zedan (11/5/1989) Wadi Habes, Egypt 3-Papaver P. argemone L. Mohamed Aid (11/4/2003) Burg El Arab, Egypt P. decaisnei Hochst & St. Ayyad et al. (21/3/1989) Mersa Matruh Siwa Road, Egypt S. Zedan (1/4/2003) Burg El Arab, Egypt P. dubium L. Ayyad et al. (21/3/1989) Mersa Matruh Siwa Road, Egypt Ayyad et al. (21/3/1989) Gara Oasis, Egypt S. Zedan (1/4/2003) Burg El Arab, Egypt P. hybridum L. A.Fakhry . (23/3/1992) Burg El Arab, Egypt S. Zedan (1/5/2001) Burg El Arab, Egypt S. Zedan (1/4/2003) Burg ElArab, Egypt L.Boulos (16/2/1979) El Hammam, Egypt T.M.Tadros (26/3/1952) Mersa Matruh, Egypt P. rhoeas L. L. Bidak (1/4/2002) Abu Seer, Egypt L. Bidak (24/4/2002) Burg El Arab, Egypt 4th .year students (11/4/1993) Burg El Arab, Egypt Ayyad (21/3/1989) Siwa, Egypt G. El Ghazaly 43 (29/2/1980) Mersa Matruh, Egypt L. Boulos (26/4/1979) Mersa Matruh, Egypt M. Rezk (10/4/1982) Noubaria, Egypt M. Rezk (21/4/1982) El Bouseily, Egypt M. Rezk (19/3/1982) Burg El Arab, Egypt Tadros (4/1949) Burg El Arab, Egypt 4-Roemeria R. hybrida (L.) DC. Ayyad (22/3/1976) Burg El Arab, Egypt L.Boulos (16/2/1979) El Hammam, Egypt L.Boulos (8/3/1979) Burg El Arab, Egypt M. Rezk (28/3/1992) Rasheed, Egypt M. Rezk (3/1993) El Bousily, Egypt G. El Ghazaly (29/2/1980) Mersa Matruh, Egypt Fumariaceae 5-Dicentra D. Formosa Walp. Gurdev Khush 22 (10/5/1959) Placer County, California 6-Fumaria F. bracteosa Pomel A. Fakhry (23/3/1992) Burg El Arab, Egypt S. Zedan (3/3/1989) Burg El Arab, Egypt R. Saad (25/3/1988) Burg El Arab, Egypt R. Saad (23/3/1988) Rasheed, Egypt Ayyad (19/3/1989) Mersa Matruh Siwa Road Ayyad (3/1992) Mersa Matruh Siwa Road Ayyad (24/3/1986) Mersa Matruh Alexandria Road Ayyad (21/3/1989) Bier nesf El Ein, Siwa, Egypt B. Bakr (2/1986) Mersa Matruh, Egypt F. densiflora DC L. Boulos (9/3/1979) Burg El Arab, Egypt L. Boulos (26/4/1979) Mariut, Egypt L. Boulos (29/2/1980) Matruh, Obayed, Egypy M. Rezk (21/4/1982) El Bousely, Egypt M. Rezk (10/4/1982) Burg El Arab, Egypt 4th year students (19/3/1982) Burg El Arab, Egypt 4th year students (19/3/1982) El Hammam, Egypt. F. judaica Boiss Tadros (5/4/1945) Smouha, Alexandria, Egypt Tadros (4/1949) Moharrem Bey, Alexandria, Egypt S. El Dareer (19/3/1999) Moharrem Bey, Alexandria, Egypt L. Boulos (5/4/1979) El Ma`amura, Alexandria, Egypt
4 Kamal W.T.
S. Kamal (16/4/1993) Mersa Matruh, Egypt F. microstachys Kral S. Zedan (12/5/1989) Agiba, Matruh, Egypt F. officinalis L. Tadros (13/3/1948) Boutonnet, Montpellier, France F. parviflora Lam Tadros (3/1945) Matruh, Egypt Hypecoaceae 7-Hypecoum H. aegyptiacum Forssk. Tadros (18/2/1945) Burg El Arab, Egypt Tadros (15/3/1945) Burg El Arab, Egypt G. El Ghazaly 375 (2/4/1982) El Omayed, Egypt S. Zedan (14/3/1988) El Omayed, Egypt I. Abdel Megid (24/1/1989) El Hammam, Egypt B. Abdel Aziz Burg El Arab, Egypt H. pendulum L. G. El Ghazaly 287 (31/3/1981) St. Katherine, Sinai, Egypt Table (2): Leaf and fruit variations within the studied taxa.
TAXA Plant Petiole Leaf Leaf Stigma Fruit Hairs Length Insertion Shape margin Arrang. Shape Length Type Eschscholzia Hairy 6 8 Radical V.F.D. Entire Basal Pointed Linear 3 3.5 Siliqua E. caespitosa E. californica Hairy 6 14 Radical F. D. Entire Basal Pointed Linear 4 5.5 Siliqua E. glyptosperma Hairy 2.5 3 Radical F. D. Entire Basal Pointed Linear 2 6 Siliqua E. lemmonii Hairy 4 6 Radical F. D. Entire Basal Pointed Linear 2 4.5 Siliqua E. lobii Hairy 3 5 Radical V.F.D. Entire Basal Pointed Linear 3 4 Siliqua E. minutifolia Hairy 1 3 Radical F. D. Entire Basal Pointed Linear 3.5 3.8 Siliqua Glaucium Hairy 0 1 Cauline pinnatifid Entire Alternate Capitate Linear 10 11 Hairy G. arabicum Siliqua G. corniculatum Hairy 0 5 Cauline Pinnatifid Entire Alternate Capitate Linear 3 6 Hairy Siliqua Papaver Hairy 3 7 Cauline Pinnatifid Entire Alternate Capitate Rect. 1 15 H.Cap./P P. argemone P. decaisnei Hairy 3 3.5 Cauline Pinnatifid Dentate Alternate Capitate Rect. 1 1.2 H.Cap./P P. dubium Hairy 2 3.5 Cauline Pinnatifid Dentate Basal Capitate Rect. 0.5 0.7 H.Cap./P P. hybridum Hairy 1 1.5 Cauline Pinnatifid Dentate Alternate Capitate Rect. 0.4 0.7 H.Caps./T P. rhoeas Hairy 1.5 3.5 Cauline Pinnatifid Dentate Alternate Capitate Rect. 1 1.2 H.Cap./P Roemeria Hairy 0 0.5 Radical F. D. Dentate Basal Pointed Linear 4 5 Hairy R. hybrida Siliqua Dicentra Glab. 1 2 Radical Pinnatifid Entire Basal Biforked Glob. 0.3 0.4 Nut D. Formosa Fumaria Glab. 1.5 2 Cauline V.F.D. Entire Alternate Biforked Glob. 0.1 0.2 Nut F. bracteosa F. densiflora Glab. 1 1.5 Cauline V.F.D. Entire Alternate Biforked Glob. 0.1 Nut. F. judaica Glab. 1.3 2 Cauline Pinnatifid Dentate Laxus Biforked Glob. 0.2 0.5 Nut F. microstachys Glab. 0 Cauline F. D. Entire Laxus Biforked Glob. 0.3 0.5 Nut F. officinalis Glab. 1 1.8 Cauline F. D. Entire Alternate Biforked Linear 0.3 0.5 Siliqua F. parviflora Glab. 1 1.5 Cauline V.F.D. Entire Alternate Biforked Linear 0.3 0.5 Siliqua. Hypecoum Glab. 2 3 Radical Pinnatifid Dentate Basal Biforked Linear 3 4 Siliqua H. aegyptiacum H. pendulum Glab. 0 Radical V.F.D. Entire Basal Pointed Linear 2 3 Siliqua Key to table 2: Glab.= Glabrous; Arrang.= arrangement; V.F.D.= very finely dissected; F. D.= finely dissected; Rect.= rectangular; Glob.= Globular; H.Caps./P= hairy capsule open by pores; H.Cap./T= hairy capsule open by teeth stigmas which ramify in a rosette like appearance over Dicentra and Hypecoum showed glabrous plants with rectangular hairy capsules. These capsules opened by both sessile or short petioled leaves and very finely pores or by teeth. Species of Fumaria have biforked dissected leaf blades. Their fruits are either nuts or stigmas, small glabrous fruits and either globular nutlets glabrous siliqua with biforked stigmas and short styles. or short linear siliquas. Hypecoum species have either These genera form a homogenous group and are pointed or biforked stigmas over very short styles and acceptable as being subfamily Fumarioideae as linear siliquas. proposed by Hoot et al. (1997). The separation of the From the above mentioned results, it is obvious that genus Hypecoum , as proposed by Hoot et al. (1999), in the three families are closely related in their another family is not supported as all the studied genera morphological criteria and their combination as one from related groups. family is more acceptable. In spite of the differences of Genera of subfamily Papaveroideae are hairy plants the stigmas, fruit shapes and types, but these differences with very long petioles, except in Glaucium species the are not enough to separate them in different families, leaves are sessile or with very short petioles. The leaf especially if the molecular data and other tools of blades are either pinnatifid, very finely or finely taxonomy showed clear similarities between them as dissected. The fruits are either glabrous or hairy siliquas mentioned above. The Studied species of Fumaria, or capsules which opened by teeth or pores and ended
5 Systematic review of the Papaveracea and status of some genera
Eschscholzia lemmonii Eschscholzia caespitosa Eschscholzia glyptosperma Eschscholzia glyptosperma
Eschscholzia lobii Eschscholzia minutiflora Glaucium arabicum Glaucium corniculatum
Papaver rhoes Papaver hybridum Papaver dubium Papaver argemone
Papaver decaisnei Roemeria hybrida Dicentra Formosa Fumaria bracteosa
Fumaria judaica Fumaria densiflora Hypecoum aegyptiacum Hypecoum parviflorum Plate (1): Photographs of selected herbarium sheets show differences in leaves and fruits. Arrows indicate to fruit.
6 Kamal W.T.
with sessile capitate or pointed stigmas with short Dargestelt Anserologischen Unterschungen in styles. These genera form another homogenous group Bereich der Rhoeadales. Plant Medic. (Stuttgard) 10 : which can be divided into two tribes; one contains the 283 297. Papaver species only and the second contain Glaucium , GERSHENZON , J., AND T.J. MABRY . 1983. Secondary Roemeria and Eschscholzia . This classification in metabolites and the higher classification of partial agreement with Hoot et al. (1997), but separate angiosperms. Nordic Journal of Botany 3: 5 34. the genus Roemeria from the genus Papaver . HEGNAUER , R. 1961. Die Gliederung der Rhoeadales The results obtained showed that genus Eschscholzia sensu Wettstein in Lichte der Inhaltsstoffe. Plant has its characteristic features which might enables it to Medic (Stuttgard) 10 : 283 297. be upgraded to the subfamily level; Eschscholzioideae ; HESLOP HARRISON , Y., AND K.P. SHIVANA . 1977. The as mentioned by Ernst (1962), Layka (1976), Heslop receptive Surface of the Angiosperm Stigma. Annals Harison and Shivana (1977), Mabry (1973), Kaderreit of Botany (London) 41 : 1233 1258. (1993), Kaderreit et al. (1994) and Bruckner (2000) but HEYWOOD , V.H. 1993. Flowering plants of the world. the species studied still have great similarities with the Andromeda Oxford Ltd. other Papaveroideae genera. HILU , K.W., T. BORSCH , AND K. MULLER ET AL . (16 CO AUTHORS ). 2003. Angiosperm phylogeny based on REFERENCES matK sequence information. American Journal of BEHNKE , H.D., AND W. BARTHLOTT . 1983. New Botany 90 : 1758 1776. Evidence from the Ultrastructural and HOOT , S.B., AND P.R. CRANE . 1995. Interfamilial Micromorphological Fields in Angiosperm relationships in the Ranunculidae based on molecular Classification. Nordic Journal of Botany 3: 43 66. systematics. Plant Systematics and Evolution BENSON , L.D. 1979. Plant Classification. Ed.2. D.C. (Supplemen) 9: 119 131. 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7 Systematic review of the Papaveracea and status of some genera
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8 Kamal W.T.
درا ا ا و و ا س ا ا ی ءـو آ ل ی ـــ ا ت، آ ا م، ا ﺱ ری ، ا ﺱ ری ، ا ا ـــ ه ا راﺱ آ ا وا راﺱ ت ا ی ا و ا س ا ا ﺏ ا ا . وﺏ دراﺱ ت رﺏ ﻥ ً ی ا ا ا ، ﺱ أﻥ اع ی ا ا ا ری وﻥ ن ی ا ا ا آ ی . وه ه ا راﺱ إ ى ﻥ ه ا ﻥ اع وه ا ﺥ ت ا ا روﺱ ﺱ دى إ ا راء ا ى ه ا ﻥ اع أم ا راء ا ﺽ ه ا ﻥ اع واﺡ . آ ا راء ا ا ی ا و ا س ا ا ﺏ ا ا وأوﺽ ع ه ا س ا أﺱ س ا ا ت ا ا وى وا و ت ا . آ و ا راﺱ ﺽ وﺽ ا ا ﺏ ا ورة وا راء ﺡ ل ا ﺏ وﺏ آ ا ا ری وا ری وا . وا ا و ﻥ أن آ ا ﻥ اع ذات ﺹ ت رﺏ ، ﺏ و د ﺏ ا ﺥ ت، و ا راء ا دى ﺏ ه ا ﻥ اع واﺡ ة . ت ا ا ر وراق رﺏ ﺡ آ ا ﻥ اع ذات أوراق ﺏ ذات أ ق و ا ل أو وه إ أوراق ری ج ة ا ق أو ا ق ﺏ دل أو ول ﺡ ل ا ق . ا ر و ا ل و آ ﻥ ﻥ ع ا د ا ودة ﺏ ات ی أو ء ﺏ ﺏ أﻥ اع ا ری ر آ ﻥ ﺏ و ا ش ا ر آ ﻥ إ ﺏ ب أو أﺱ ن .
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